Even in the highest class of the Mollusca, namely the Cephalopoda or cuttle-fishes, in which the sexes are separate, secondary sexual characters of the kind which we are here considering, do not, as far as I can discover, occur. This is a surprising circumstance, as these animals possess highly-developed sense-organs and have considerable mental powers, as will be admitted by every one who has watched their artful endeavours to escape from an enemy.411 Certain Cephalopoda, however, are characterised by one extraordinary sexual character, namely, that the male element collects within one of the arms or tentacles, which is then cast off, and, clinging by its sucking-discs to the female, lives for a time an independent life. So completely does the cast-off arm resemble a separate animal, that it was described by Cuvier as a parasitic worm under the name 326of Hectocotyle. But this marvellous structure may be classed as a primary rather than as a secondary sexual character.
Although with the Mollusca sexual selection does not seem to have come into play; yet many univalve and bivalve shells, such as volutes, cones, scallops, &c., are beautifully coloured and shaped. The colours do not appear in most cases to be of any use as a protection; they are probably the direct result, as in the lowest classes, of the nature of the tissues; the patterns and the sculpture of the shell depending on its manner of growth. The amount of light seems to a certain extent to be influential; for although, as repeatedly stated by Mr. Gwyn Jeffreys, the shells of some species living at a profound depth are brightly coloured, yet we generally see the lower surfaces and the parts covered by the mantle less highly coloured than the upper and exposed surfaces.412 In some cases, as with shells living amongst corals or brightly-tinted sea-weeds, the bright colours may serve as a protection. But many of the nudibranch mollusca, or sea-slugs, are as beautifully coloured as any shells, as may be seen in Messrs. Alder and Hancock’s magnificent work; and from information kindly given me by Mr. Hancock, it is extremely doubtful whether these colours usually serve as a protection. With some species this may be the case, as with one which lives on the green leaves of algæ, and is itself bright-green. But many brightly-coloured, white or otherwise conspicuous species, do not seek concealment; whilst again some equally conspicuous species, as well as other dull-coloured kinds, live under stones and in 327dark recesses. So that with these nudibranch molluscs, colour apparently does not stand in any close relation to the nature of the places which they inhabit.
These naked sea-slugs are hermaphrodites, yet they pair together, as do land-snails, many of which have extremely pretty shells. It is conceivable that two hermaphrodites, attracted by each others’ greater beauty, might unite and leave offspring which would inherit their parents’ greater beauty. But with such lowly-organised creatures this is extremely improbable. Nor is it at all obvious how the offspring from the more beautiful pairs of hermaphrodites would have any advantage, so as to increase in numbers, over the offspring of the less beautiful, unless indeed vigour and beauty generally coincided. We have not here a number of males becoming mature before the females, and the more beautiful ones selected by the more vigorous females. If, indeed, brilliant colours were beneficial to an hermaphrodite animal in relation to its general habits of life, the more brightly-tinted individuals would succeed best and would increase in number; but this would be a case of natural and not of sexual selection.
Sub-kingdom of the Vermes or Annulosa: Class, Annelida (or Sea-worms).—In this class, although the sexes (when separate) sometimes differ from each other in characters of such importance that they have been placed under distinct genera or even families, yet the differences do not seem of the kind which can be safely attributed to sexual selection. These animals, like those in the preceding classes, apparently stand too low in the scale, for the individuals of either sex to exert any choice in selecting a partner, or for the individuals of the same sex to struggle together in rivalry.
328Sub-kingdom of the Arthropoda: Class, Crustacea.—In this great class we first meet with undoubted secondary sexual characters, often developed in a remarkable manner. Unfortunately the habits of crustaceans are very imperfectly known, and we cannot explain the uses of many structures peculiar to one sex. With the lower parasitic species the males are of small size, and they alone are furnished with perfect swimming-legs, antennæ and sense-organs; the females being destitute of these organs, with their bodies often consisting of a mere distorted mass. But these extraordinary Fig. 3. Labidocera Darwinii,
(from Lubbock).
a. Part of right-hand anterior antenna of male, forming a prehensile organ.
b. Posterior pair of thoracic legs of male.
c. Ditto of female. differences between the two sexes are no doubt related to their widely different habits of life, and consequently do not concern us. In various crustaceans, belonging to distinct families, the anterior antennæ are furnished with peculiar thread-like bodies, which are believed to act as smelling-organs, and these are much more numerous in the males than in the females. As the males, without any unusual development of their olfactory organs, would almost certainly be able sooner or later to find the females, the increased number of the smelling-threads has probably been acquired through sexual selection, by the better provided males having been the most successful in finding partners and in leaving offspring. Fritz Müller has described a remarkable dimorphic species of Tanais, in which the male is represented by two distinct forms, never graduating into each other. In the one form the male is furnished with more numerous smelling-threads, and in the other form with more powerful and more elongated chelæ or pincers which serve to hold the female. Fritz Müller suggests that these differences between the two male forms of the same species must have originated in certain individuals having varied in the number of the smelling-threads, whilst other individuals varied in the shape and size of329 their chelæ; so that of the former, those which were best able to find the female, and of the latter, those which were best able to hold her when found, have left the greater number of progeny to inherit their respective advantages.413
In some of the lower crustaceans, the right-hand anterior antenna of the male differs greatly in structure from the left-hand one, the latter resembling in its simple tapering joints the antennæ of the female. In the male the modified antenna is either swollen in the middle or angularly bent, or converted (fig. 3) into an elegant, and sometimes wonderfully complex, prehensile organ.414 It serves, as I hear from Sir J. Lubbock, to hold the female, and for this same purpose one of the two posterior legs (b) on the same side of the body is converted into a forceps. In another family the inferior or posterior antennæ are “curiously zigzagged” in the males alone.
330
Fig. 4. Anterior part of body of Callianassa (from Milne-Edwards), showing the unequal and differently-constructed right and left-hand chelæ of the male.
N.B.—The artist by mistake has reversed the drawing, and made the left-hand chela the largest.
Fig. 5. Fig. 6.
Fig. 5. Second leg of male Orchestia Tucuratinga (from Fritz Müller).
Fig. 6. Ditto of female.
In the higher crustaceans the anterior legs form a pair of chelæ or pincers, and these are generally larger in the male than in the female. In many species the chelæ on the opposite sides of the body are of unequal size, the right-hand one being, as I am informed by Mr. C. Spence Bate, generally, though not invariably, the largest. This inequality is often much greater in the male than in the female. The two chelæ also often differ in structure (figs. 4 and 5), the smaller one resembling those of the female. What advantage331 is gained by their inequality in size on the opposite sides of the body, and by the inequality being much greater in the male than in the female; and why, when they are of equal size, both are often much larger in the male than in the female, is not known. The chelæ are sometimes of such length and size that they cannot possibly be used, as I hear from Mr. Spence Bate, for carrying food to the mouth. In the males of certain freshwater prawns (Palæmon) the right leg is actually longer than the
whole body.415 It is probable that the great size of one leg with its chelæ may aid the male in fighting with his rivals; but this use will not account for their inequality in the female on the opposite sides of the body. In Gelasimus, according to a statement quoted by Milne-Edwards,416 the male and female live in the same burrow, which is worth notice, as shewing that they pair, and the male closes the mouth of the burrow with one of its chelæ, which is enormously developed; so that here it indirectly serves as a means of defence. Their main use, however, probably is to seize and to secure the female, and this in some instances, as with Gammarus, is known to be the case. The sexes, however, of the common shore-crab (Carcinus mænas), as Mr. Spence Bate informs me, unite directly after the female has moulted her hard shell, and when she is so soft that she would be injured if seized by the strong pincers of the male; but as she is caught and carried about by the male previously to the act of moulting, she could then be seized with impunity.
Fritz Müller states that certain species of Melita are 332distinguished from all other amphipods by the females having “the coxal lamellæ of the penultimate pair of feet produced into hook-like processes, of which the males lay hold with the hands of the first pair.” The development of these hook-like processes probably resulted from those females which were the most securely held during the act of reproduction, having left the largest number of offspring. Another Brazilian amphipod (Orchestia Darwinii, fig. 7) is described by Fritz Müller, as presenting a case of dimorphism, like that of Tanais; for there are two male forms, which differ in the structure of their chelæ.417 As chelæ of either shape would certainly have sufficed to hold the female, for both are now used for this purpose, the two male forms probably originated, by some having varied in one manner and some in another; both forms having derived certain special, but nearly equal advantages, from their differently shaped organs.
It is not known that male crustaceans fight together for the possession of the females, but this is probable; for with most animals when the male is larger than the female, he seems to have acquired his greater size by having conquered during many generations other males. Now Mr. Spence Bate informs me that in most of the crustacean orders, especially in the highest or the Brachyura, the male is larger than the female; the parasitic genera, however, in which the sexes follow different habits of life, and most of the Entomostraca must be excepted. The chelæ of many crustaceans are weapons well adapted for fighting. Thus a Devil-crab (Portunus puber) was seen by a son of Mr. Bate fighting with a Carcinus mænas, and the latter was soon thrown on its back, and had every limb torn from its body. 333 When several males of a Brazilian Gelasimus, a species furnished with immense pincers, were placed together by Fritz Müller in a glass vessel, they mutilated and killed each other. Mr. Bate put a large male Carcinus mænas into a pan of water, inhabited by a female paired with a smaller male; the latter was soon dispossessed, but, as Mr. Bate adds, “if they fought, the victory334 was a bloodless one, for I saw no wounds.” This same naturalist separated a male sand-skipper (so common on our sea-shores), Gammarus marinus, from its female, both of which were imprisoned in the same vessel with many individuals of the same species. The female being thus divorced joined her comrades. After an interval the male was again put into the same vessel and he then, after swimming about for a time, dashed into the crowd, and without any fighting at once took away his wife. This fact shews that in the Amphipoda, an order low in the scale, the males and females recognise each other, and are mutually attached.
Fig. 7. Orchestia Darwinii (from Fritz Müller), showing the differently-constructed chelæ of the two male forms.
The mental powers of the Crustacea are probably higher than might have been expected. Any one who has tried to catch one of the shore-crabs, so numerous on many tropical coasts, will have perceived how wary and alert they are. There is a large crab (Birgos latro), found on coral islands, which makes at the bottom of a deep burrow a thick bed of the picked fibres of the cocoa-nut. It feeds on the fallen fruit of this tree by tearing off the husk, fibre by fibre; and it always begins at that end where the three eye-like depressions are situated. It then breaks through one of these eyes by hammering with its heavy front pincers, and turning round, extracts the albuminous core with its narrow posterior pincers. But these actions are probably instinctive, so that they would be performed as well by a young as by an old animal. The following case, however, can hardly be so considered: a trustworthy naturalist, Mr. Gardner,418 whilst watching a shore-crab (Gelasimus) making its burrow, 335threw some shells towards the hole. One rolled in, and three other shells remained within a few inches of the mouth. In about five minutes the crab brought out the shell which had fallen in, and carried it away to the distance of a foot; it then saw the three other shells lying near, and evidently thinking that they might likewise roll in, carried them to the spot where it had laid the first. It would, I think, be difficult to distinguish this act from one performed by man by the aid of reason.
With respect to colour which so often differs in the two sexes of animals belonging to the higher classes, Mr. Spence Bate does not know of any well-marked instances with our British crustaceans. In some cases, however, the male and female differ slightly in tint, but Mr. Bate thinks not more than may be accounted for by their different habits of life, such as by the male wandering more about and being thus more exposed to the light. In a curious Bornean crab, which inhabits sponges, Mr. Bate could always distinguish the sexes by the male not having the epidermis so much rubbed off. Dr. Power tried to distinguish by colour the sexes of the species which inhabit the Mauritius, but always failed, except with one species of Squilla, probably the S. stylifera, the male of which is described as being “of a beautiful blueish-green,” with some of the appendages cherry-red, whilst the female is clouded with brown and grey, “with the red about her much less vivid than in the male.”419 In this case, we may suspect the agency of sexual selection. With Saphirina (an oceanic genus of Entomostraca, and therefore low in the scale) the males are furnished with 336minute shields or cell-like bodies, which exhibit beautiful changing colours; these being absent in the females, and in the case of one species in both sexes.420 It would, however, be extremely rash to conclude that these curious organs serve merely to attract the females. In the female of a Brazilian species of Gelasimus, the whole body, as I am informed by Fritz Müller, is of a nearly uniform greyish-brown. In the male the posterior part of the cephalo-thorax is pure white, with the anterior part of a rich green, shading into dark brown; and it is remarkable that these colours are liable to change in the course of a few minutes—the white becoming dirty grey or even black, the green “losing much of its brilliancy.” The males apparently are much more numerous than the females. It deserves especial notice that they do not acquire their bright colours until they become mature. They differ also from the females in the larger size of their chelæ. In some species of the genus, probably in all, the sexes pair and inhabit the same burrow. They are also, as we have seen, highly intelligent animals. From these various considerations it seems highly probable that the male in this species has become gaily ornamented in order to attract or excite the female.
It has just been stated that the male Gelasimus does not acquire his conspicuous colours until mature and nearly ready to breed. This seems the general rule in the whole class with the many remarkable differences in structure between the two sexes. We shall hereafter find the same law prevailing throughout the great sub-kingdom of the Vertebrata, and in all cases it is eminently distinctive of characters which have been 337acquired through sexual selection. Fritz Müller421 gives some striking instances of this law; thus the male sand-hopper (Orchestia) does not acquire his large claspers, which are very differently constructed from those of the female, until nearly full-grown; whilst young his claspers resemble those of the female. Thus, again, the male Brachyscelus possesses, like all other amphipods, a pair of posterior antennæ; the female, and this is a most extraordinary circumstance, is destitute of them, and
so is the male as long as he remains immature.
Class, Arachnida (Spiders).—The males are often darker, but sometimes lighter than the females, as may be seen in Mr. Blackwall’s magnificent work.422 In some species the sexes differ conspicuously from each other in colour; thus the female of Sparassus smaragdulus is dullish-green; whilst the adult male has the abdomen of a fine yellow, with three longitudinal stripes of rich red. In some species of Thomisus the two sexes closely resemble each other; in others they differ much; thus in T. citreus the legs and body of the female are pale-yellow or green, whilst the front legs of the male are reddish-brown: in T. floricolens, the legs of the female are pale-green, those of the male being ringed in a conspicuous manner with various tints. Numerous analogous cases could be given in the genera Epeira, Nephila, Philodromus, Theridion, Linyphia, &c. It is often difficult to say which of the two sexes departs most from the ordinary coloration of the genus to which the species belong; but Mr. Blackwall 338thinks that, as a general rule, it is the male. Both sexes whilst young, as I am informed by the same author, usually resemble each other; and both often undergo great changes in colour during their successive moults before arriving at maturity. In other cases the male alone appears to change colour. Thus the male of the above-mentioned brightly-coloured Sparassus at first resembles the female and acquires his peculiar tints only when nearly adult. Spiders are possessed of acute senses, and exhibit much intelligence. The females often shew, as is well known, the strongest affection for their eggs, which they carry about enveloped in a silken web. On the whole it appears probable that well-marked differences in colour between the sexes have generally resulted from sexual selection, either on the male or female side. But doubts may be entertained on this head from the extreme variability in colour of some species, for instance of Theridion lineatum, the sexes of which differ when adult; this great variability indicates that their colours have not been subjected to any form of selection.